Table Of Content) 5
The Wilson Journal ofOrnithology 118(4):537-546, 2006
NESTING SUCCESS OF GRASSLAND AND SAVANNA BIRDS ON
3
RECLAIMED SURFACE COAL MINES OF THE MIDWESTERN
UNITED STATES
EDWARD W. GALLIGAN,1’ TRAVIS L. DeVAULT,24 AND STEVEN L. LIMA1
—
ABSTRACT. Reclaimed surface coal mines in southwestern Indiana support many grassland and shrub/
savanna bird species of conservation concern. We examined the nesting success of birds on these reclaimed
mines to assess whether such “unnatural” places represent productive breeding habitats for such species. We
established eight study sites on two large, grassland-dominated mines in southwestern Indiana and classified
them into three categories (open grassland, shrub/savanna, and a mixture ofgrassland and shrub/savanna)based
on broad vegetation and landscape characteristics. During the 1999 and 2000 breeding seasons, we found and
monitored 911 nests of 31 species. Daily nest survival for the most commonly monitored grassland species
ranged from 0.903 (Dickcissel, Spiza americana) to 0.961 (Grasshopper Sparrow, Ammodramus savannarum).
Daily survival estimates for the dominant shrub/savanna nesting species ranged from 0.932 (Brown Thrasher,
Toxostoma rufum) to 0.982 (Willow Flycatcher, Empidonaxtrailin'). Vegetation and landscapeeffects on nesting
success were minimal, and only Eastern Meadowlarks (Sturnella magna) showed a clear time-of-season effect,
with greaternesting success in the firsthalfofthe breeding season. Rates ofBrown-headedCowbird (Molothrus
ater parasitism were only 2.1% forgrassland species and 12.0% for shrub/savanna species. The nesting success
ofbirds on reclaimed mine sites was comparable to that in other habitats, indicating that reclaimed habitats on
surface mines do not necessarily represent reproductive traps for birds. Received 1 August 2005, accepted 10
April 2006.
Several bird species have benefited in re- contain scattered trees (from plantings and
cent decades from the reclamation of surface natural succession) that approximate the struc-
coal mines in the midwestem United States ture of savanna habitat to a substantial degree
(Bajema et al. 2001, DeVault et al. 2002, In- (Scott et al. 2002, Scott and Lima 2004). Ac-
gold 2002). The Surface Mining Reclamation cordingly, these reclaimed mines harbor sev-
Act of 1977 and earlier laws led (perhaps un- eral savanna bird species (DeVault et al. 2002)
intentionally) to mine reclamation techniques of conservation concern (Davis et al. 2000,
that favored the production ofgrasslands rath- Hunter et al. 2001).
er than forested habitats (Brothers 1990), re- The size of reclaimed mines in the mid-
sulting in hundreds of km2 of newly created westem United States—is one oftheir most im-
grasslands. These “mine grasslands” harbor a portant characteristics several exceed 2,000
diverse assemblage of grassland birds, many ha (Bajema and Lima 2001, Ingold 2002).
of which are of management concern at state Many grassland bird species appear to be
and federal levels. Recent studies in south- “area sensitive” in that usually they are found
western Indiana, covering 19 reclaimedmines,
only in grassland fragments of a given size or
suggest that populations ofkey grassland bird
greater (Herkert 1994, Walk and Warner 1999,
species, such as Grasshopper {Ammodramus
savannarum) and Henslow’s (A. henslowii) Winter and Faaborg 1999; but see Horn et al.
2000, Johnson and Igl 2001). Most studies
DspeaVraruolwts,etareal.qu2i0t0e2)la.rgReec(lBaaijmeemda emtinale.s20a0l1s,o suggest that grasslands >50-100 ha should
contain a full complement of grassland pas-
serines. Virtually all grasslands on reclaimed
Sta1tDeeUpnti,v.o,fETecrorleogHyauatned, OIrNga4n7i8s0m9a,lUBSiAo.logy, Indiana mines in southwestern Indiana are >100 ha
2Dept, of Forestry and Natural Resources, Purdue (Bajema and Lima 2001). Furthermore, small
Univ., West Lafayette, IN 47907, USA. grassland size may be associated with poor
3Current address: Louisville Metro Health Dept., nesting success, reflecting the close proximity
400 E. Gray St., Louisville, KY 40202, USA. of habitat edge, which can lead to greater
4Current address: U.S. Dept, of Agriculture, Wild-
life Services, National Wildlife Research Center, 5757 predator densities (Winter et al. 2000, Herkert
Sneller Rd., Brewerton, NY 13029, USA. et al. 2003) and greaterrates ofBrown-headed
5Corresponding author; e-mail: [email protected] Cowbird {Molothrusater) parasitism (Johnson
537
538 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 118, No. 4, December2006
and Temple 1990). These effects of habitat counties and four were established at the Uni-
size are similar to those documented for many versal Mine (39° 36' N, 87° 28' W; 3,450 ha)
forest-nesting passerines (e.g., Donovan et al. in southern Vermillion County. The Chinook
1995, Robinson et al. 1995). sites ranged in size from 39 to 67 ha, whereas
Even though large reclaimed coal mines in the Universal sites were smaller (12 to 38 ha)
the Midwest harbor a variety ofbreeding bird due to constraints imposed by cattle and hay-
species, they are decidedly unnatural places in ing operations. Chinook Mine comprised 61%
terms of vegetation (Scott and Lima 2004). undisturbed grassland and 18% hayfields; the
%
Hence, it is conceivable that reclaimed mines remaining 21 comprised relatively even per-
function as giant ecological “traps” that di- centages of wetlands, row crops, and forests
vert breeding birds away from more produc- (Bajema and Lima 2001). Universal Mine was
tive habitats {sensu Gates and Gysel 1978). 33% undisturbed grassland and 43% hayfields
Even though grassland birds can breed suc- and cattle pastures, with the remaining 24%
cessfully in non-native grasslands (e.g., Warn- split about evenly between forest and lakes/
er 1994, Best et al. 1997, Kershner and Bol- wetlands (Bajema and Lima 2001).
linger 1998, Robb et al. 1998, Ingold 2002, Study sites were chosen to represent the
Monroe and Ritchison 2005), the possibility range of grassland-dominated habitats found
that they represent ecological traps is not triv- in the reclaimed surface coal mines of south-
ial. For example, reclaimed midwestern mines western Indiana. Two study sites (one in each
often are dominated by tall fescue (Festuca mine) were classified as “open grassland.”
arundinacea; Scottet al. 2002, Scottand Lima We defined open grassland sites as relatively
2004), which often is infected with a symbi- undisturbed areas (no mowing for >2 years,
otic fungal endophyte (Neotyphodium coeno- usually many more) that were dominated by
phialum). Such infected fescue is associated grasses (>95%, by area), with some forbs and
with declines in plant diversity and lowered very few saplings, trees, or shrubs (Scott et al.
reproductive success of herbivores (vertebrate 2002). Open grasslands represented the most
and invertebrate; Clay and Holah 1999). Tall abundant habitat type found on most re-
fescue might reduce insect production and claimed surface mines (Bajema and Lima
render reclaimed mine grasslands into poor 2001). Nests found in these open sites were,
m
breeding habitat. Although tall fescue also on average, 760 from the nearest mature
may affect the breeding prospects of savanna forest habitat, with many nests well over
m
bird species, they might be less affected than 1,000 from forest.
their grassland counterparts. Three study areas were classified as “shrub/
There are few data available for assessing savanna” sites (one at Chinook Mine and two
whether birds inhabiting reclaimed surface at Universal Mine). We defined shrub/savanna
mines are nesting successfully. Thus, our goal sites as predominantly grassy habitats with
in this study was to investigate patterns ofavi- many scattered young trees (4-8 m high, gen-
an nesting success within reclaimed surface erally open canopy) and shrubs, often repre-
coal mines, with the larger goal of evaluating senting a transition zone between grassland
whether reclaimed mines provide productive and forested areas. Small groves of trees also
breeding habitats for grassland and savanna were associated with small wetland areas.
birds. Black locusts (Robinia pseudoacacia) domi-
nated in shrub/savanna sites, although signif-
METHODS
icant numbers of oaks Quercus spp.), eastern
— (
Study sites. Our work spanned the 1999 cottonwoods (Populus deltoides), and mature
and 2000 breeding seasons. In both years, autumn olives (Elaeagnus umbellata) were
field work began in late April and continued found in some areas. “Shrubby” species in-
through the 1st week of August. Study sites cluded young saplings of these tree species,
were established at two large reclaimed sur- along with hawthorn (Crataegus spp.) and
face coal mines in west-central Indiana within multiflora rose bushes (Rosa multiflora).
30 km of the city of Terre Haute. Four sites Shrub/savanna sites were adjacent to mature
were established at the Chinook Mine (39° forest (and hence were mainly on the edges of
28' N, 87° 13' W; 2,000 ha) in Clay and Vigo the reclaimed mines). The average distance
-
Galligan et al. • NESTING SUCCESS ON RECLAIMED SURFACE MINES 539
between nests found on shrub/savanna sites nearest forest edge, and distance to the nearest
and the forest edge was 240 m. Shrub/savanna tree (>1 m high). We used GPS units to re-
sites contained significant (30-60%, by area) cord the location of all nests and to delineate
open grassland habitat. nearby forested areas.
—
Finally, we designated the remaining three Data analyses. We estimated the daily
study sites (two at Chinook Mine and one at probability of nest survival (DNS) for each
Universal Mine) as “mixed” sites. Mixed species according to the Mayfield method
sites were defined as mostly open grassland (Mayfield 1961, 1975). We assumed that any
habitat with a few areas of significant shrub/ relevant nesting event (e.g., hatching, failure,
savanna habitat. In general, these sites were fledging) occurred at the midpoint of the in-
70-80% open grassland. The average distance terval between nest visits. A nest was consid-
between nests found on mixed sites and the ered successful when it fledged one or more
nearest mature forest habitat was 430 m. young (Mayfield 1961, 1975).
—
Nest location and monitoring. Nest Our analyses were limited primarily to uni-
searches were conducted daily in 1999 and variate tests of vegetation, landscape, and
2000 from early morning until early afternoon temporal variable effects on DNS or the fate
by a team ofthree to five field workers. Nests of individual nests (success or failure). We
were detected by (1) rope dragging, (2) fol- tested for interactions only for study site and
lowing adults that were carrying food and time of season. We compared DNS estimates
nesting material, and (3) systematic searches across categorical variables (i.e., among years,
of likely nesting sites (Martin and Geupel sites, and different habitat types) by using
1993). During the 2000 field season, we also CONTRAST (Hines and Sauer 1989). CON-
used a thermographic imager to aid in nest TRAST uses a generalized x2 statistic that al-
detection (Galligan et al. 2003). lows multiple comparisons of survival rates
When a nest was located, a small colored from different time periods or study areas
flag was placed 10 m to the north of it and a (Sauer and Williams 1989). We compared
small piece of colored tape was tied to vege- DNS among years and sites for all species list-
tation 5 m south of the nest (Picozzi 1975, ed in Table 1. Because we found large num-
Walk 2001). The accurate alignment of flag, bers of Field Sparrow (Spizella pusilla), Am-
tape, and nest allowed workers to relocate modramus spp. (Henslow’s and Grasshopper
nests quickly with minimal disturbance. Spe- sparrows, combined), Dickcissel Spiza amer-
(
cies associated with each nest were identified, icana), Red-winged Blackbird (Agelaius
and nests were checked only every 3 to 4 days phoeniceus), and Eastern Meadowlark (Stur
to minimize disturbance (Bart 1977). During nella magna nests, we were able to examine
)
each nest check, we recorded the presence or DNS trends within breeding seasons (compar-
absence of adults, the number of eggs or ing DNS between the first and second halves
chicks, and, if appropriate, the developmental of the breeding seasons) and between habitat
stage of the chicks. We also recorded indica- types for these species. We used logistic re-
tors of nestling mortality or cowbird parasit- gression, with the fate of individual nests
ism. (failure or success) as the dependent variable,
—
Vegetation and landscape variables. We to evaluate the effects of various continuous
gathered basic information on the physical re- landscape and vegetation variables on nesting
lationships between nests, the surrounding success (SPSS, Norusis 1993). Our analyses
vegetation, and major landscape features; were applied primarily to habitat types (open,
however, we limited analyses of these vari- mixed, and shrub/savanna) because they were
ables to grassland bird species, whose nests distinctly different from the surrounding land-
were located in greater numbers than savanna scape characteristics. For a given habitat type,
species. For each nest, we recorded height we limited our analyses to those species for
above ground, species and height of the veg- which we had adequate numbers of nest-days
etation in which it was placed, dominant veg- (see grassland species listed in Table 1). The
m
etation and vegetation height within 1 of effects of various factors on nest survival
the nest, litter depth at the nest, percent cover were analyzed individually, except as noted.
m
of litter within 1 ofthe nest, distance to the Results are presented as means and standard
540 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 118, No. 4, December2006
TABLE 1. Daily nest survival (DNS) for bird species inhabiting grassland and shrub/savanna on reclaimed
coal mines in Indiana during 1999 and 2000.
n(no.successful) NDa DNS SE
Grassland species
Grasshopper Sparrow 41 (26) 383 0.961 0.010
Henslow’s Sparrow 21 (9) 236 0.949 0.014
Field Sparrow 90 (36) 629 0.919 0.011
Dickcissel 47 (11) 369 0.903 0.016
Eastern Meadowlark 129 (58) 1450 0.951 0.006
Red-winged Blackbird 264 (74) 2439 0.923 0.005
Shrub/Savanna species
Mourning Dove 62 (33) 816 0.962 0.007
Willow Flycatcher 30 (22) 440 0.982 0.006
American Robin 33 (12) 313 0.933 0.014
Brown Thrasher 31 (14) 251 0.932 0.016
Yellow Warbler 21 (13) 272 0.971 0.010
aNumberofnest-daysobserved.
errors; the level of significance was set at Daily probability of nest survival: overall
—
0.05. estimates. The overall estimates of DNS (all
data pooled) showed considerable interspecif-
RESULTS ic variation. Among grassland species (Table
During our 2-year study, we found 911 ac- 1), we estimated relatively high rates of DNS
tive nests of 31 species. Of these nests, 465 (near 0.950) for Grasshopper Sparrows, Hen-
and 446 were found at the Chinook and Uni- slow’s Sparrows, and Eastern Meadowlarks.
versal mines, respectively. Red-winged Black- Conversely, we estimated DNS of <0.925 for
birds, Eastern Meadowlarks, Field Sparrows, Dickcissels (the lowest: 0.903), Field Spar-
Dickcissels, Grasshopper Sparrows, and Hen- rows, and Red-winged Blackbirds. Among sa-
slow’s Sparrows were (in that order) the best vanna species. Willow Flycatchers and Yellow
represented grassland birds (Table 1). Nests of Warblers (Dendroica petechia) experienced
other grassland species, such as those ofRing- the greatest DNS (0.982 and 0.971, respec-
necked Pheasants Phasianus colchicus tively); we also estimated a high DNS for
Sedge Wrens Cisto(thorus platensis an)d, Mourning Doves (0.962), and our DNS esti-
( ),
Bobolinks Dolichonyx oryzivorus were too mate for Brown Thrasher was the lowest
( ),
few in number for analyses, as these species (0.932).
are relatively rare on the reclaimed surface Predation accounted for the vast majority of
mines (DeVault et al. 2002). Among the nest failures. In most cases, we could only
shrub/savanna species, nests of Mourning guess at the identity of the predators involved
Doves Zenaida macroura), Willow Flycatch- because many predators do not leave conclu-
(
ers (Empidonax traillii), American Robins sive evidence of their identities at depredated
(Turdus migratorius), and Brown Thrashers nests (Thompson et al. 1999, Maier and
('Toxostoma rufum) were found most frequent- DeGraaf 2000, Pietz and Granfors 2000, Bur-
ly (Table 1). The nests of other savanna spe- hans et al. 2002). However, many snakes were
cies were located in numbers too small for encountered during nest searches, mainly rac-
analyses, including those of Eastern Kingbird ers (Coluber constrictor) and black rat snakes
(Tyrannus tyrannus), Bell’s Vireo (Vireo bel- (Elaphe obsoleta obsoleta); we also encoun-
lii), Song Sparrow (Melospiza melodia), Blue tered smallernumbers ofgarter snakes (Tham-
Grosbeak Passerina caerulea), Indigo Bun- nophis spp.) and prairie kingsnakes (Lampro-
(
ting (Passerina cyanea), Orchard Oriole (Ic- peltis calligaster). Snakes were observed con-
terus spurius), and American Goldfinch (Car- suming eggs or chicks on two occasions. Only
duelis tristis). Red-winged Blackbirds appeared to sufferany
Galligan etal. • NESTING SUCCESS ON RECLAIMED SURFACE MINES 541
TABLE 2. Daily nest survival (DNS) for grassland birds, by site type (open grassland, mixed, and shrub/
savanna), on reclaimed coal mines in Indianaduring 1999 and 2000; x2 statistics weredeterminedusingprogram
CONTRAST.
Species Habitattype n DNS SE
Red-winged Blackbird Open grassland 97 0.914 0.010
Mixed 154 0.923 0.007
Shrub/savanna 21 0.949 0.014
X2 = 4.13, df = 2, P = 0.13
Eastern Meadowlark Open grassland 62 0.939 0.010
Mixed 46 0.962 0.008
Shrub/savanna 23 0.974 0.010
X2 = 6.39, df = 2, P = 0.04
Dickcissel Open grassland 23 0.903 0.022
Mixed 15 0.916 0.024
Shrub/savanna 9 0.871 0.043
X2 = 0.86, df = 2, P = 0.65
Field Sparrow Open grassland 0. 33 0.933 0.017
Mixed 25 0.938 0.016
Shrub/savanna 34 0.879 0.024
II 45- o df = 2, P = 0.11
Ammodramus spp. Open grassland 25 0.977 0.009
Mixed 20 0.943 0.017
Shrub/savanna 17 0.928 0.023
X2 = 6.04, df = 2, P = 0.05
weather-induced mortality (nests blown over 926 ± 0.011, respectively; x2 — 12.33, df =
during severe thunderstorms), and then only 1, P < 0.001).
early in the 1999 breeding season. There were Significant differences in DNS also were
no indications of significant nutritional stress observed across habitat types (Table 2). DNS
among any nestlings. for Eastern Meadowlarks was greatest in
—
Effects of time and site. DNS estimates shrub/savanna habitat (0.974) and lowest in
(all sites pooled) did not differ between years the open habitats (0.939; x2 = 6.39, df = 2,
(sXa2v:aanlnlaPspveacliueesse>xce0p.1t0B)rfoorwnanTyhrgarsahsesrla(nxd2o=r wPas= h0i.g0h4e1r).inDNthSe oinfcAremamsiondgrlyamoupsensphaabrirtoawtss
5.70, df = 1, P = 0.017). Brown Thrasher (X2 = 6.04, df = 1, P = 0.050). For Field
DNS was very low in 1999 (0.895 ± 0.027), Sparrows, our DNS estimates tended to be
but was much greater in 2000 (0.969 ± lower in the shrub/savanna habitats (x2 =
0.015). For American Robin, there was a sim- 4.508, df = 1, P = 0.11). DNS for Dickcissels
ilar across-year trend (x2 — 3.22, df = 1, P = also was lowest in the shrub/savanna habitat
0.072) in DNS, which increased from 0.885 (0.871), but not significantly so. Logistic re-
± 0.034 to 0.951 ± 0.014. gression analyses of these data produced very
We found a significant time-of-season ef- similar results, indicating no significant inter-
fect only for Eastern Meadowlarks; in both actions between habitat type and time of sea-
years, our estimate oftheir DNS was substan- son, for any of the species listed in Table 2
tially greater during the first halfofthe breed- (Wald x2 tests: all P > 0.50). DNS did not
ing season than in the second half. In 1999, differ between mines (x2: all P > 0.10; pool-
their DNS decreased from 0.974 ± 0.008 to ing data across all study sites within a given
0.919 ± 0.016 (x2 = 9.45, df = 1, P = 0.005) mine) for any species listed in Table 1.
and, in 2000, from 0.966 ± 0.009 to 0.934 ± Effects of vegetation and landscape vari-
0.014 (x2 = 3.70, df = 1, P = 0.051). When ables.—Our analyses indicated few significant
the data were pooled across years, DNS in the associations between DNS and vegetation or
first and second half of the breeding season landscape features. However. DNS forEastern
differed substantially (0.970 ± 0.006 versus Meadowlarks increased with distance to forest
542 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 118, No. 4, December2006
bin t=he0s.0h1r9u,b/sWaavladnnxa2si=tes3(.l9o5g,istdifc r=egr1e,ssPion=: hosTtAsBpLecEies3.wasBrionwfnr-ehqueeandtedonCorewcbliaridmepdarcaosailtimsimneosf
0.047). Increasing nest height also was asso- in Indiana during 1999 and 2000.
ciated with lower DNS forField Sparrows, but
only in open grassland habitats (b = —0.487, Species3 n ParaNsoit.ized %
Wald x2 = 4.22, df = 1, P = 0.040); DNS Grassland
was lower for nests in low shrubs than for
Sedge Wren l 0 0.0
those on the ground. For Red-winged Black- Red-winged Blackbird 263 4 1.5
birds, height of vegetation in which the nest Bobolink 1 0 0.0
was placed was positively associated with Eastern Meadowlark 131 0 0.0
nesting success, but only in the mixed habitat Dickcissel 47 2 4.1
type (b = 0.051, Wald x2 = 6.42, df = 1, P Field Sparrow 93 6 6.4
= 0.011). Finally, for Grasshopper Sparrows, Grasshopper Sparrow 41 1 2.4
m Henslow’s Sparrow 21 0 0.0
height of the dominant vegetation within 1 Total 607 13 2.1
of the nest was positively associated with
Shrub/Savanna
nesting success, but only when the data were
pooled across all habitat types (b = 0.465, Eastern Kingbird 9 1 11.1
Wald x2 = 4.14, df = 1, P = 0.046). It is WBeillll’sowViFrleyocatcher 306 01 106..70
notable that tall fescue (either as the vegeta- Yellow Warbler 21 3 14.3
tion in which the nest was placed or as the Orchard Oriole 10 4 40.0
m
dominant vegetation within 1 of the nest) Blue Grosbeak 6 2 33.3
was not significantly associated with the DNS Indigo Bunting 4 0 0.0
of any focal species. American Goldfinch 6 0 0.0
Brood parasitism.—Relatively low rates of Song Sparrow 8 1 12.5
Total 100 12 12.0
brood parasitism by Brown-headed Cowbirds
were observed during our 2-year study. Over- 3Known egg rejectors (e.g., American Robins, Brown Thrashers)and
unsuitablecowbirdhosts(e.g..MourningDoves)werenotincluded.
all, only 2.1% of grassland bird nests were
parasitized by cowbirds (Table 3). Field Spar-
rows were the most heavily parasitized relatively high rates of DNS, whereas Amer-
(6.4%), whereas we observed no parasitism on ican Robins and Brown Thrashers experienced
Henslow’s Sparrows or Eastern Meadowlarks. relatively low rates of DNS. There were no
Furthermore, of the 263 Red-winged Black- significant differences in DNS across the two
bird nests that we found, only four were par- mines studied, despite the fact that these
asitized. Shrub/savanna species as a group (in- mines encompass the range of land-use pat-
cluding all species monitored) suffered a terns found within mines (Bajema and Lima
greater frequency ofbrood parasitism (12.0%; 2001). There also were few significant differ-
Table 3). Ofthe savanna species. Orchard Ori- ences in DNS across the two breeding sea-
oles and Blue Grosbeaks were most heavily sons, despite the fact that the first season
parasitized. (1999) was relatively hot and dry, and the sec-
ond season (2000) was cool and wet (only
DISCUSSION
Brown Thrashers and American Robins had
—
Daily nest survival. Overall estimates of markedly greater DNS in 2000 than 1999).
DNS varied considerably across species. In Thus, the general patterns apparent in Table 1
general, shrub/savanna birds experienced may be representative of the long-term situa-
greater rates of DNS than grassland birds (Ta- tions faced by birds on the reclaimed mines
ble 1). Among grassland birds. Eastern Mead- of southwestern Indiana.
owlarks, Grasshopper Sparrows, and Hen- Ultimately, the variation that we observed
slow’s Sparrows experienced relatively high in DNS was due to variation in nest predation,
rates of DNS, whereas Dickcissels, Field the primary cause of nest failure. Among
Sparrows, and Red-winged Blackbirds expe- grassland birds, it appears that open-cup,
rienced lower rates of DNS. Among shrub/sa- above-ground nesters, such as Field Sparrows,
vanna species. Mourning Doves, Willow Fly- Dickcissels, and Red-winged Blackbirds, suf-
catchers, and Yellow Warblers experienced fered greater predation rates than ground-nest-
Galligan et al. • NESTING SUCCESS ON RECLAIMED SURFACE MINES 543
ing species (Eastern Meadowlarks and Am- shrub/savanna habitat than elsewhere (Table
modramus sparrows; Table 1). During both 2).
field seasons, we estimated greater DNS for Across studies, a consistent picture of the
Eastern Meadowlarks during the first half of effects of vegetation and landscape variables
the breeding season than during the second on nesting success of many grassland species
half. This time-of-season effect may reflect has yet to emerge. For example, Johnson and
the fact that Eastern Meadowlarks began nest- Temple (1990) observed increased nest pre-
ing in April before snakes became fully ac- dation for grassland passerines when their
tive. No other temporal patterns in DNS were nests were located near wooded edges. Winter
apparent among other grassland species. et al. (2000) found that, for artificial nests,
Open-cup nesting was not uniformly asso- fragment size and vegetation characteristics
ciated with greater rates of nest predation, be- were better predictors of survival than dis-
cause all shrub/savanna species in this study tance to habitat edge; however, Henslow’s
are open-cup nesters, and many experienced Sparrow nests placed within 50 m of an edge
high rates of DNS (Table 1). The relatively were not as successful as those at greater dis-
low rate of nesting success among American tances from forest edge. For Dickcissels, dis-
Robins and Brown Thrashers was due to ex- tance to habitat edge also appeared to have
tremely high levels of nest predation during little effect on daily survival in prairie habitats
1999 (which was not observed in 2000). Why (Hughes et al. 1999, Winter et al. 2000). Bur-
only these two species experienced different hans et al. (2002) observed that Field Spar-
levels of predation across years is not clear; rows nesting in old fields had greater success
however, because robins and thrashers nested when nest height was >3 m above ground;
in very similar sites in the shrub/savanna hab- however. Best (1978) suggested that Field
itat (interior portions of larger trees), they Sparrows were more successful when nests
likely experienced the same change in the were near the ground or in relatively tall veg-
predatory environment across years. etation. Pribil (1998) did not detect a relation-
Significant associations between DNS and ship between nest success and vegetation fea-
various vegetation and landscape-level fea- tures for Red-winged Blackbirds.
—
tures were few, and provided relatively little Brood parasitism. Brood parasitism was
insight into the predation processes that influ- minimal in our focal species, especially when
enced DNS. We note, however, that for many compared with the high frequency of parasit-
species we located too few nests for our anal- ism reported in midwestem forest fragments
yses to detect subtle effects. Regardless, the (e.g., Robinson et al. 1995). For grassland
significant increase in D—NS with increasing birds, only 2.1% of nests were parasitized.
distance from the forest exhibited only in The frequency of parasitism for Red-winged
Eastern—Meadowlarks in the shrub/savanna Blackbirds at our reclaimed surface coal
habitat was consistent with the results of mines (1.5%) markedly contrasts with the par-
other studies (e.g., Johnson and Temple 1990) asitism frequency of >30% for this species in
that implicated forest-edge predators as major other habitats and areas to the west of our
agents of nest failure (recall that our shrub/ study sites (Yasukawa and Searcy 1995, Clot-
savanna sites were adjacent to forested habi- felter and Yasukawa 1999). Kershner (2001)
tat). The lack ofan effect ofdistance-to-forest and Walk (2001) reported similarly low fre-
in the open grassland and mixed study sites quency of parasitism for grassland birds nest-
may reflect the relative isolation ofthese sites ing in restored prairies in nearby eastern Illi-
from forested habitat (c/. Paton 1994). The nois (see also Robinson and Herkert 1997,
relatively high rates ofDNS forAmmodramus Kershner and Bollinger 1998). Perhaps the
sparrows in the open grassland habitats (Table frequency of grassland bird parasitism is gen-
2) also may reflect the isolation from forested erally greater well to the west of Indiana
habitat. Nevertheless, there was no association (Johnson and Temple 1990, Zimmerman
between distance-to-forest and DNS for any 1993, Davis 2003; but see Winter 1999, Win-
other species in the shrub-savanna sites. Fur- ter et al. 2004). In any case, the low frequency
thermore, the overall nesting success of East- of cowbird parasitism for grassland birds of
ern Meadowlarks was actually greater in the western Indiana and eastern Illinois supports
544 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 118, No. 4, December2006
TABLE4. Daily nest survival forgrassland birds on reclaimed coal mines in Indianaduring 1999 and 2000
was similar to that recorded at other midwestern grassland sites.
Species KanfsiaesldsC®RP P(rMaOir)ieb NBWigRO(aIkNs)C (egIgo,wnaesCtRliPngfisetladgse)d prRaiersiteor(eILd)e Remcilnaeisme(dINc)ofal
— — —
Red-winged Blackbird — — 0.943—, 0.916 0.954 0.923
Eastern Meadowlark 0.940 — 0.953 0.951
Dickcissel 0.—922 0.—940 0.951,—0.874 0.941 0.903
Field Sparrow — 0.—919 0.955 0.919
Grasshopper Sparrow — 0.930 0.957,—0.937 0.—913 0.961
Henslow’s Sparrow 0.950 0.947 0.949
aHughesetal.(1999).bWinterandFaaborg(1999),cRobbetal.(1998),dPattersonandBest(1996),eKershner(2001)andWalk(2001);fthisstudy.
the idea that cowbirds in the eastern United essentially identical to that observed in our
States focus on forested habitats (Hahn and reclaimed mine sites (Table 4). Dickcissels
Hatfield 1995). Indeed, Brown-headed Cow- and Field Sparrows experienced greater nest
bird is among the rarest passerine species in- success at the Illinois sites than at our sites,
habiting reclaimed coal mines in Indiana whereas Grasshopper Sparrows experienced
(DeVault et al. 2002). greater success at our mine sites (few Hen-
Shrub/savanna species underwent greater slow’s Sparrow nests were found at the Illi-
rates ofparasitism than grassland species (Ta- nois site). Similar to what we found in our
ble 3), but it was still much lower than that study. Red-winged Blackbirds in Conserva-
typically observed in forested habitats across tion Reserve Program (CPR) fields of Iowa
Indiana and Illinois (e.g., Robinson et al. experienced poor to mediocre nesting success,
1995). Among the focal savanna species (Ta- Dickcissels experienced low success (with
ble 1), only Yellow Warblers were parasitized very low survival in the nestling stage), and
to a substantial degree (Table 3). Note, how- Grasshopper Sparrows had relatively high
ever, that three of our focal savanna species rates of success (Patterson and Best 1996).
are either inappropriate cowbird hosts Dickcissels also may not be doing well in
(Mourning Doves) or egg rejectors (American Kansas or Missouri CRP fields (Hughes et al.
Robins and Brown Thrashers). Parasitism ap-
1999, Winter and Faaborg 1999). Nesting suc-
peared to be greater for some non-focal sa-
cess of Field and Henslow’s sparrows at the
vanna species (e.g.. Orchard Orioles and Blue Big Oaks National Wildlife Refuge (formerly
Grosbeaks; Table 3), but we found too few the Jefferson Proving Ground) in southeastern
nests to reach a conclusion concerning their
We Indiana is virtually identical to that of birds
susceptibility to parasitism. suspect that
nesting on reclaimed surface coal mines
greater rates of cowbird parasitism in our
shrub/savanna sites reflected their proximity (Robb et al. 1998). Furthermore, survival es-
to forested habitat (Hahn and Hatfield 1995). timates for Henslow’s Sparrows across the
Conservation implications—Our data sug- three relevant studies (Robb et al. 1998, Win-
gest that reclaimed surface co.al mines are no ter and Faaborg 1999; this study) were re-
more likely to represent reproductive traps markably similar and relatively high, indicat-
than are other habitats studied to date. We ing that this species is probably doing reason-
base this view on a comparison of our results ably well where it is still nesting. Similarly,
with those from comparable studies across the Monroe and Ritchison (2005) reported com-
midwestern United States. DNS within re- parable levels of nesting success for Hen-
claimed coal mine grasslands at our study slow’s Sparrows on reclaimed mines and un-
sites is broadly comparable to that in other mined grasslands in western Kentucky, and
midwestern grasslands (Table 4). The most suggested that reclaiming surface mines could
comparable study is one that took place in help stabilize the population decline of Hen-
large blocks of restored prairies in nearby slow’s Sparrows. We suspect that similar con-
eastern Illinois (Kershner 2001, Walk 2001). clusions also could be drawn for some savan-
DNS of Eastern Meadowlarks in Illinois was na species on reclaimed mines, but compara-
Galligan etal. • NESTING SUCCESS ON RECLAIMED SURFACE MINES 545
ble data are not yet available with which to Best, L. B. 1978. Field Sparrow reproductive success
make analogous comparisons. and nesting ecology. Auk 95:9-22.
Reclaimed mines of the Midwest provide a Best, L. B., H. Campa, III, K. E. Kemp, R. J. Robel,
puenciiqaulelyopfpoorrtutnhietymainnaagvieamnenctonsoerfvagtriaosns,laensd- nMS.e.stRRi..ngRWyianinnC,teRJr.PstAe.fiineSl.advs1i9da9gn7ed.,cHBr.iorpPd.laWanedbeuknisnd,atnhJcre.e,Maiandnd-d
birds. Many of the reclaimed mines are west: a regional approach. Wildlife Society Bul-
>2,000 ha, larger than most (if not all) re- letin 25:864-877.
maining prairie fragments in Indiana and Il- Brothers, T. S. 1990. Surface-mine grasslands. Geo-
linois, and contain large populations ofseveral graphical Review 80:209-225.
bDierVdauslpetceitesal.of20c0o2n,cIenrgnol(dBa20j0e2m)a.eTtheal.ne2s0t0i1n,g Burhaaatnnsdso,Jn.gDbF.iaraEdb.,onersDgt..s.D2Je0o0au2rr.bnaoFlrancot,foWrFis.ladRfl.fiefTcehtiMonamgnppsarogened,amteiInIoItn,
success of key species (e.g., Henslow’s Spar- 66:240-249.
rows and Grasshopper Sparrows) at these re- Clay, K. andJ. Holah. 1999. Fungalendophytesym-
claimed mines is comparable with that in non- biosis and plant diversity in successional fields.
mined grassland habitats. A feature that Science 285:1742-1744.
should make reclaimed midwestern surface Clotfelter, E. D. and K. Yasukawa. 1999. Impact
of brood parasitism by Brown-headed Cowbirds
coal mines attractive from a management per- on Red-winged Blackbird reproductive success.
spective is that they are usually owned by a Condor 101:105-114.
single entity. Furthermore, most reclaimed Davis, M. A., D. W. Peterson, P. B. Reich, M. Cro-
mines are typically not very productive as ag- zier, T. Query, E. Mitchell, J. Huntington,and
ricultural areas. These factors combined make P. Bazakas. 2000. Restoring savanna using fire:
possible the acquisition or management of impact on the breeding bird community. Restora-
large grassland-dominated habitats. Few such tion Ecology 8:30-40.
Davis, S. K. 2003. Nesting ecology of mixed-grass
opportunities currently exist in the eastern prairie songbirds in southern Saskatchewan. Wil-
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ACKNOWLEDGMENTS Lima. 2002. Breeding bird communities of re-
claimed coal-mine grasslands in the American
The U.S. Fish and Wildlife Service, the U.S. Geo- Midwest. Journal of Field Ornithology 73:268-
logical Survey, and the Ohio River Valley Ecosystem 275.
Group provided financial support for this project. We Donovan, T. M., F. R. Thompson, III,J. Faaborg,and
are grateful to the following individuals for allowing J. R. Probst. 1995. Reproductive success of mi-
us full access to reclaimed mine properties: A. Eicher gratory birds in habitat sources and sinks. Con-
and S. McGarvie of Peabody Coal, R. Ronk of the servation Biology 9:1380-1395.
Indiana Department ofNatural Resources, M. Krieger Galligan, E. W., G. S. Bakken,andS. L. Lima. 2003.
atUniversal Mine, andL. Nelsonofthe MidwestCoal Using a thermographic imager to find nests of
Company. We alsothankG. S. Bakken, M. T. Jackson, grassland birds. Wildlife Society Bulletin 31:865-
and P. E. Scott for valuable help and advice. Special 869.
thanks go to R. Gushee, J. Mozingo, C. Roever, B. Gates, J. E. and L. W. Gysel. 1978. Avian nest dis-
Thomas, and A. Worthingtonfortheircompetentfield- persion and fledging success in field-forest eco-
work and keen nest-finding abilities. We are grateful tones. Ecology 59:871-883.
to M. D. Carey and two anonymous reviewers for Hahn, D. C. and J. S. Hatfield. 1995. Parasitism at
helpful comments that improved this manuscript. the landscape scale: cowbirds prefer forest. Con-
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